thoughts about saami prehistory

THOUGHTS ABOUT SAAMI PREHISTORY John Weinstock, University of Texas

Introduction Is there a clearer picture today than there was when K. B. Wiklund viewed the vestiges of the Komsa culture as “an aboriginal glacial and interglacial Lapp culture”?1 Speculation began in earnest in the mid-19th century when Sven Nilsson claimed that the Lapps were the original inhabitants of Scandinavia. This idea was soon opposed by Gustaf von Düben who asserted that the Lapps had migrated into Scandinavia relatively recently from the East. Gustaf Hallström and Wiklund were later supporters of von Düben’s views; but everything changed in the late 1920s when Anders Nummedal discovered Stone Age remains along the arctic coast of Norway, the so-called Komsa culture. Since Wiklund’s death in 1934 speculation about the origins of the Saami has continued unabated (cf. among many Christian Carpelan, Milton Nuñez, Markku Niskanen, Kalevi Wiik). Our knowledge has improved significantly since then: archaeological excavations have turned up much new information; historical linguistics provides a better idea of the interrelationship of the Finno-Ugric languages; and population genetic studies, especially, have become much more sophisticated. Yet each of the data sets from archaeology, paleoenvironmental science, linguistics and genetics are in a sense incompatible, for they often belong to widely differing time perspectives and geographical areas. It is like trying to finish a jigsaw puzzle lacking most of the pieces. Moreover, each of the data types has its own issues: archaeological sites and finds can be dated quite accurately through C-14 calculations, but there are often significant gaps in time and place. Historical linguistics extrapolates back to hypothetical language families for which there is no physical evidence at all, and the construction of trees for daughter languages (linguistic phylogeny) is rife with controversy. Analysis of inherited genetic material, as Lars Ivar Hansen and Bjørnar Olsen write, can “reflect degrees of affinity, contact and common origin,

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Manker 1952: 42. Wiklund’s death in 1934 prevented him from completing his investigation, but his theory was published posthumously, e.g. in Wiklund 1947; cf. Wiklund 1947: 3–17.

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while differences may indicate isolation and historical distance.”2 Genetic analysis extrapolates back from contemporary blood and urine samples to relationships between peoples thousands of years ago; hence, conclusions reached are fraught with difficulties such as time scale, sampling choices, selection criteria, and more. DNA analysis of ancient bones might be of more help in the future. As Colin Renfrew, referring to Luigi Luca Cavalli-Sforza’s maps, aptly puts it: […] the difficulty is that these maps, reflecting the current situation (since they are based on samples derived from living populations) represent a palimpsest of all the historical processes and events which have taken place since the first population of Europe by Homo sapiens.3

Thus, one must be very cautious in making inferences. It is also difficult to draw reliable conclusions based on only one type of data. If corroboration can be found between two or more data types we are on safer ground. Since this article focuses on the Saami and Saami ethnicity it important to make clear what we mean by ethnicity. Following the lead of Fredrik Barth and his introduction to Ethnic Groups and Boundaries Hansen and Olsen see Saami ethnic identity developing through consciousness of their (the Saami’s) “cultural dissimilarity from others” that came about through contact,4 e.g. between trapping communities in northern Fennoscandia and Russian metal-producing communities to the east. They date the development of Saami ethnicity toward the end of the last millennium BC. This means two things: first, much of the DNA analysis takes us back to the Paleolithic era, thousands of years before there was any Saami ethnicity. In other words, the peoples mentioned here may be ancestors of the Saami, Finns, Norwegians, Swedes, etc. Secondly, Hansen and Olsen caution that “the easy [read wrong] conclusion is that ‘ethnicity is in the blood,’ and that Saami ethnicity should be viewed as a physical and biological package that can be traced tens of thousands of years back in time.”5

Early Settlement Northern Europe opened up to settlement after glacial ice began to retreat ca. 19 500 BC. Until then glaciers covering Fennoscandia and the Alps were separated by a thin corridor, with refuges in the Franco-Cantabrian (Iberian) area to the west

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Hansen & Olsen (2004) 2007: ch. 2.7. Renfrew 2000: 18. 4 Hansen & Olsen (2004) 2007: ch. 2.6. 5 Hansen & Olsen (2004) 2007: ch. 2.7. 3

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and on the South Russian Plain to the east; but there was little interaction nor exchange of genes between the two areas while the ice was at its most widespread. As the glaciers retreated human groups began to move into newly ice-free areas. These were not migrations in the usual sense, in that the ice retreated only a few kilometers per generation.6 To the west Late Paleolithic reindeer hunters moved north, settling the British Isles and the North European Plain. Their most significant genetic feature was the maternal DNA (mtDNA) haplogroup V.7 Some of these hunters reached the coast of Northern Norway by 9 000 BC and were known as the Komsa culture. The eastern complex occupying an area south and east of the ice from Eastern Poland to Northwest Russia settled much of Northeastern Europe. Forest and fresh water resources were important for them and their most significant genetic feature was the paternal Y-chromosomal C allele.8 They may have been speakers of Uralic languages.9 However, the traditional view among Uralic/Finno-Ugric historical linguists is that speakers of the language ancestral to Uralic lived in the Volga region about 6 000–4 000 BC, implying that some of them migrated from there to Fennoscandia.10 That would mean there were no Uralic/Finno-Ugric languages spoken in Fennoscandia until long after the ice had melted. However, in 1995 Pekka Sammallahti proposed that there “may well have been genetic and linguistic ancestors of the modern Finns and Saami immediately after the ice-cap receded in Scandinavia.”11 Pavel Dolukhanov presented several arguments that supported this so-called continuity theory: “archaeological records show a considerable cultural continuity […] from the Upper Palaeolithic to the ethnographically recognizable Finns.” He added that “Uralic languages are extremely archaic: there is hardly any doubt that they predate the spread of I-E speech.” The vocabulary too reflects an ancient mode of life.12

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Nuñez 2002: 157. Niskanen 2002: 145. A haplogroup is a grouping of mtDNAs with mutational events defined by SNPs (single-nucleotide polymorphisms). V is pronounced “vee.” 8 Niskanen 2002: 145. Different alleles produce variation in inherited characteristics such as hair color or blood type. 9 There is some dispute as to whether the South Russian Plain was a Uralic homeland or whether it was settled by Proto-Indo-Europeans during the Mesolithic. Niskanen suggests that with warmer times seasonal migrations between hunting territories became longer and that peoples speaking Proto-IndoIranian dialects moved into more southerly areas vacated by speakers of Proto-Uralic dialects (Niskanen 2002: 145). 10 Campbell 1997. 11 Sammallahti 1995. Others have proposed this, e.g. Kivikoski 1961. 12 Dolukhanov 1996: 45. 7

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The inhabitants of the southeastern shore of the Baltic were members of the Late Upper Paleolithic Swiderian culture 10 800–9 700 BP.13 The Swiderian was succeeded by the Early Mesolithic Post-Swiderian cultures such as Maglemose in the west to Kunda (in Estonia) and Veretye in the east that emerged ca. 8 700 BC. Various groups were arrayed near the edge of the ice sheet, from future Germanic and Baltic people in the southwest to future Ugrians in the northeast with future Finns and (South) Saami in between. The groups to the southwest may have been speakers of some early form of Indo-European or all of these groups may have spoken some version of Uralic; but there was little genetic differentiation between them. The first inhabitants of Finland were from the Kunda culture to the south in Estonia and from the Veretye culture to the east in Karelia and they arrived ca. 8 500 BC14 or 8 500–8 000 cal BC / 9 500–9 000 BP.15 Settlers reached northernmost Finland by 7 500 cal BC16 / 8 000 BC.17 Further west there is evidence of the Maglemose culture – similar to the Kunda – in Southern Scandinavia.

Agriculture and Indo-European Languages According to Niskanen the earliest farming in the East Baltic region goes back to the Corded Ware (Battle Axe) culture 3 200–2 500 cal BC.18 The introduction of agriculture to Finland came around 4 500–4 000 BP.19 Renfrew suggested that this change in subsistence pattern was connected to the spread of Indo-European languages from the Near East to Europe – Southern and Central Europeans are closer genetically to Near Eastern groups (Renfrew 1992). Prior to the arrival of agriculture in Fennoscandia the inhabitants who had come from the southeast were primarily speakers of Finno-Ugric or Uralic tongues. The spread of farming into Northern Europe was slower and took place mainly through cultural and, eventually, linguistic rather than demic diffusion and, hence, there was not as much influence from southern genes.20 When farming arrived in the north along with

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Swiderian culture takes its name from the Swidr point; it is located in the sand dune areas of Poland, Lithuania. 14 Niskanen 2002: 146. 15 Nuñez 2002: 164. 16 Nuñez 2002: 164. 17 Niskanen 2002: 146. 18 Niskanen 2002: 147. 19 Nuñez 2002: 168. 20 Zvelebil and Zvelebil 1988; cf. also Karlsson et al. 2006, Richards et al. 2000 and Richards et al. 1996. Summaries of the bold face references can be viewed at the end of the paper.

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Indo-European languages the peoples remained largely the same, adopting Germanic, Baltic or Slavic languages, or they kept their Finno-Ugric tongues. Were Indo-European languages spoken in Scandinavia before agriculture arrived? The most likely answer is no.21 Of the various theories on the evolution of the Indo-European languages Renfrew’s above would seem to have fewer problems than competing theories such as the Kurgan hypothesis and the Paleolithic continuity theory.22 One question is that of language trees and how divergences can be dated. A recent computational phylogenetic analysis based on 87 I-E languages and 2449 cognate sets dated the split of the Germanic and Italic branches to between 5 500–1 750 BP when the Germanic languages became a separate branch in Northern Germany or Southern Scandinavia.23 The more likely earlier date would bring Proto-Germanic tongues to Scandinavia about when the agricultural idea crossed the Baltic and long after the Uralic/Finno-Ugric languages were there.24 Sammallahti reconstructed Proto-Uralic vocabulary at 4 500 BC in an area from northeastern Central Europe in the southwest to the Ural Mountains in the east.25 A relatively large number of I-E words were borrowed by various versions of the Uralic languages over time, but it was not until Proto-FinnoVolgaic26 spoken around 3 000 BC that the first words were borrowed from PreGermanic/Germanic/Baltic. At this point in time early forms of the Finno-Ugric languages had been spoken in Fennoscandia for thousands of years.

Population Genetic Analysis I – Saami Origins The Saami have a heterogeneous genetic origin and “are regarded as extreme genetic outliers among European populations.”27 Two mtDNA haplogroups, V and

21

For a contrary view cf. Appelbaum and Appelbaum who argue on the basis of Torroni et al. 1998 that I-E groups “arrived from the east and some time later the major Finno-Ugric migration-carrying the markers found on many Saami Y chromosomes-began.” Yet, there has been considerable phylogenetic analysis in the past ten years that speaks against the Appelbaums’ views. 22 The Kurgan hypothesis seems to have lost favor recently and the paleolithic continuity theory has not achieved widespread support. 23 Gray & Atkinson 2003. 24 The Germanic languages are a conundrum in attempts to come up with a genealogy for the I-E language family. Recent work in computational phylogeny suggests that at one time Germanic was closely related to the Balto-Slavic branch but then grew closer to the Celtic and Italic branches. Cf. work by Donald Ringe et al. at http://www.cs.rice.edu/~nakhleh/CPHL/. 25 Sammallahti 1998: 118. 26 Sammallahti 1998: 121 f. 27 Tambets et al. 2004.

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Map 1. Possible entry routes of the predominant Saami maternal lineages into Fennoscandia. Broken lines indicate that the exact place of origin/route of spread of the haplogroup is unsolved/not indicated (Tambets et al. 2004: 677).

U5b1b1 (the so-called Saami motif) account for ca. 80% of the Saami gene pool.28 V is common in Europe with age estimates of 16 000–8 000 BP from west to east; that is, it likely originated in the northern Iberian peninsula or southwestern France (the Franco-Cantabrian glacial refuge area) at the time of the Younger Dryas29 and spread east from there.30 U5b1b1 has a frequency of 47.6% among Saami, much lower frequencies among Finns and Karelians and is otherwise almost absent in Western Europe. Directional lines on maps 1–2 suggest plausible paths that mtDNA and Y-chromosomal lineages may have taken to Fennoscandia. The Y-chromosomal haplogroups most significant in the Saami are N3 and R1a with a combined frequency of ca. 60%, both with eastern origin. Yet the Saami’s maternally inherited mtDNA as well as their paternally inherited Y-chromosomal variation are primarily European. The large genetic separation of the Saami from other Europeans has been explained by a relatively limited and constant population size together with a long period of isolation and genetic drift. This manifests itself as linkage disequilibrium (LD).31 Ikegaya et al. used the JCV (JC virus) genotype to study Saami and Finnish origins.32 They confirm that the Saami and Finns are mainly European on the maternal side but that the Saami and not the Finns were influenced by a founder 28

Torroni et al. 1998. The Younger Dryas was the period of the “big freeze” ca. 12 700–11 500 BP. 30 Cf. also Torroni et al. 2001. 31 Cf. Johansson et al. 2004, Tambets et al. 2004 and Kaessmann et al. 2002. 32 Ikegaya et al. 2005. 29

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Map 2. Possible entry routes of the predominant Saami paternal lineages into Fennoscandia. Broken lines indicate that the exact place of origin/route of spread of the haplogroup is unsolved/not indicated. NB: what seems to be a J in the map is actually an I with a dot below (Tambets et al. 2004: 677).

effect or bottleneck phenomenon.33 This could well be the period of isolation on the North Atlantic coast. On the paternal side they infer multiple founding populations for both ethnic groups. A study by Achilli et al. confirms the FrancoCantabrian refuge as the source of the late-glacial expansion of hunter-gatherers into Northern Europe.34 These Late Paleolithic hunters, ancestors of the Komsa culture, crossed the dry North Sea bottom and reached the northernmost coast of Norway by 9 000 cal BC / 9 800 BP.35 We do not know what language they might have spoken, though it was unlikely to have been a Uralic tongue. Bergman et al. argue that the deglaciation of Fennoscandia proceeded more rapidly than once thought.36 They discovered the earliest Mesolithic site in Sweden up to that point in time; it is located north of the 60th parallel and dated at 8600 BP. They suggest that it was most likely populated by Komsa people from the north who left the coast in search of new resources.37 As mentioned above, FinnoUgrian settlers reached northernmost Finland by 7 500 cal BC / 8 000 BC where they encountered those moving south and southeast from Komsa areas; the latter 33

A bottleneck is a significant reduction in population size. A founder effect is a migration of a relatively small group of people from their home to establish a new population elsewhere followed by isolation and rapid population growth. 34 Achilli et al. 2005. The Saami and the Berbers who came from the same refuge share subclade U5b1b that contains the “Saami motif.” 35 Nuñez 2002: 162. 36 Bergman et al. 2004. 37 This is exactly what Wiklund expected new archaeological finds to show (Manker 1952: 42).

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more likely adopted the Finno-Ugrian/Uralic tongue from those they came in contact with who were better equipped to exploit the resources of the inland areas. They are among the ancestors of the (North) Saami. Complicating the picture, the mtDNA haplogroups show different frequencies among Saami groups with the Southern Swedish Saami being outliers.38 Interestingly, haplogroup Z is not found in Norwegian Saami nor Continental European and only at 1% for Northern Swedish Saami; however, it is found at higher frequencies in Finnish Saami (5%) and Southern Swedish Saami (6%). Ingman and Gyllensten see this as a contribution to the Saami gene pool from the Volga-Ural region of Russia, “a migration from Eastern Europe” going back only to 2 700 BP. The date is well within the period of contact with the Ananjino culture when there was close interaction between these metal workers and northern hunting communities. Rather than as a migration Hansen and Olsen write that “it may be explained within the transactional framework […] where social and economic transactions were firmed by marriage alliances and kinship ties.”39

Population Genetic Analysis II – Finnish Origins The origin of the Finns presents problems not unrelated to the origin of the Saami. The occurrence of several diseases among Finns that are rare elsewhere has spurred numerous genetic analyses. Sajantila et al. found that there was little Ychromosomal diversity in the Finnish population relative to other European populations and they attributed this to a male bottleneck or founder effect.40 They also found that mitochondrial mutations (decreases in genetic diversity) accumulated after the bottleneck. In other words, relatively few men and women contributed to the gene pool of today’s Finnish population. They suggested that the bottleneck occurred ca. 4 000 BP when populations using agriculture arrived in Finland. Possibly a small number of I-E speaking Corded Ware people were linguistically assimilated by the indigenous Uralic-speaking people (i.e., the Saami). However, it is unlikely that a culture with more complex social organization would be linguistically assimilated by an indigenous group. Niskanen found it more likely that the Corded Ware people arriving in Finland ca. 4 500–4 000 BP already spoke a Proto-Baltic-Finnish language and that they did not differ genetically from those hunter-gatherers already there.41 And, as mentioned above, the introduction of 38

Ingman and Gyllensten 2006. Hansen and Olsen (2004) 2007: ch. 2.7 40 Sajantila et al. 1996. 41 Niskanen 2002: 147. 39

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agriculture was not demic, i.e. it was primarily the idea of agriculture that crossed over to Fennoscandia without a significant contribution of genes from the south. Kittles et al. examined Y-chromosomal data from nine Finnish provinces and they were able to confirm a dual-origins hypothesis, i.e. that there were two founder populations.42 Haplotype A/49 is most common in the northeast near the Lake Ladoga region but has no counterpart in other Scandinavian populations. Haplotype B/69 is common in the south and is observed in other European populations. On their map Kittles et al. indicated that A/49 entered Finland from the east whereas A/69 came into Finland from the south and west. Furthermore, A/49 predated B/69 by about 2 000 years in Finland. This suggested to the authors two genetically distinct waves of male settlers, the latter arriving with the wave of agriculture/agriculturalists around 4 500–4 000 BP. Lahermo et al. studied males from sixteen Eurasian ethnic groups including the Finns and two Saami groups (Skolt and Inari).43 They found reduced variation due to bottlenecks in the male lineages of the Finns and the Saami or possibly due to drift because of small effective male population over a long period of time. Raitio et al. genotyped five Y-chromosomal microsatellite markers in Finns and Saami.44 Their study only included Inari, Kola and Skolt Saami. They concluded that the most common Y-chromosomal haplotypes are shared between Finns and Saami and that there were two Y-chromosomal founding lineages in both populations. Summing up then, the Saami and Finns are quite distinct genetically based on mtDNA sequences and autosomal markers but similar based on Y-chromosome markers. Furthermore, multiple founder effects in both populations suggest an infusion of genes from the south along with the arrival of farming ca. 4 500–4 000 BP and later possibly from the east through contact with the Ananjino culture ca. 2 700 BP.

Conclusions How are we to read the myriad of phylogenetic and phylogeographic data? The first observation is that there is a great deal of diversity within the Saami genetic constellation (see maps 3–4). Even though none of the authors mentioned in the summaries surveyed the entire spectrum of Saami groups and lifestyles, there is enough data to reveal clear phylogeographic patterns throughout Europe and, in

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Kittles et al. 1998. Lahermo et al. 1999. 44 Raitio et al. 2001. 43

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Map 3. Variation in the motif U5b1b1 between four Saami groups.

particular, among the Saami. Consider the following map which is just one of many that could be drawn to demonstrate such geographic patterns: Might there be a scale running northwest to southeast with those Saami in areas to the south and southeast having a greater input of paternal and maternal genes from the southeast/east? The mtDNA figures on map 3 support this: the Saami motif U5b1b1 e.g. varies from 56.8% in the northwest to 23.9% to the south.45 The Y-chromosomal data present a similar picture: three haplogroups – N3, I and R1a – make up more than 80% of the Saami Y-chromosomal gene pool. In map 4 the great variation in these three haplogroups can be seen, with the Finnish Saami as outliers.46

45 46

Cf. also Tambets et al. 2004. Cf. Tambets et al. 2004, Rootsi et al. 2004, Rosser et al. 2000

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Map 4. Variation of the three haplogroups N3, I and R1a between three Saami groups.

Peoples tend to influence one another through transactions near and far, trade and intermarriage; thereby they also influence one another genetically. Raitio et al., as mentioned above, found that the Finns and Saami (Inari, Kola, Skolt) share the most common Y-chromosomal haplotypes though with regional differences.47 Finally, there have been many contributions to today’s Saami gene pool, or even to the Saami gene pool at the end of the first millennium BC when Saami ethnicity came into existence. Since this input came from just about every direction and corner of Europe it makes no sense to talk about a place of origin for the Saami, unless one means Fennoscandia where these genes arrived along with the many forebears of the Saami. Saami culture today and in the recent past exhibits striking heterogeneity and that would also seem to be the case for the origins of the Saami people genetically and archaeologically, and probably linguistically as well.

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Raitio et al. 2001.

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Summaries Brief summaries of 31 articles from 1995 to the present (2007) involving, directly or indirectly, phylogenetic analysis of the Saami. Articles may be accessed by searching lead author and first few words of title. Achilli et al. 2004 “The Molecular Dissection of mtDNA […]”: analysis of mtDNA haplogroup H: Franco-Cantabrian refuge major source of European gene pool – accords well with earlier finding of haplogroup V in age and spread. Achilli et al. 2005 “Saami and Berbers […]”: U haplogroup: Saami and Berbers share a young subclade, U5b1b (U5b1b1 is Saami motif) – confirms Franco-Cantabrian refuge as source of late-glacial expansion. Delghandi et al. 1997 “Saami Mitochondrial DNA […]”: two major maternal lineage clusters in Norwegian Saami: Lineage 1 (more diverse) from 76% in Kautokeino to Coast Northern and Lineage 2 (less diverse) from 22.6% in Kautokeino to 39.7% in Karasjok (but 9.1% in Finnish Inarii Saami) – authors suggest two independent migration waves of small founding populations with Lineage 1 older. Finnilä et al. 2000 “Phylogenetic Network of the mtDNA Haplogroup U […]”: polymorphisms of mtDNA of 22 Finns subclustered in U5 ➪ U restricted in variation in Finland – can be extrapolated to entire population. Hedman et al. 2004 “Analysis of 16 Y STR […]”: Finnish subpopulations in the eastern parts of the country lack genetic variation ➪ small male founder population and bottleneck indicated by reduced Y-chromosomal heterogeneity compared to Europeans on the average. Hedman et al. 2006 “Finnish mitochondrial DNA […]”: mtDNA haplogroups H (40%) and U (27.5%) predominate among group of 200 Finnish males [65.5% of the U is the Saami motif] ➪ diversity comparable with rest of Europe – Hedman et al. 2004 found minimal Ychromosomal heterogeneity vs. rest of Europe – higher female migration rate could explain diversity difference. Ikegaya et al. 2005 “Genetic Diversity […]”: population history of Saami and Finns has been inferred, based on studies using mtDNA and Y chromosome as markers: studies based on JCV genotype and mtDNA suggest both populations mostly of Caucasoid origin, but studies based on Y chromosome suggest Asian contribution to both groups – EU-a1 for 87% of Saami (vs. Finns 33%) ➪ single founding population – curiously EU-a1 also found in Siberian/Inuit and NE Japanese ➪ migration from Central Asia in two directions. Ingman and Gyllensten 2006 “A recent genetic link […]”: majority of mtDNA diversity in northern Swedish, Norwegian and Finnish Saami accounted for by haplogroups V and U5b1b1; southern Swedish Saami have other haplogroups and frequency distribution similar to Continental European population – comes from recent admixture with Swedish population – divergence time for V is 7 600 BP and for U5b1b1 is 5 500 BP among Saami and 6 600 BP among Saami and Finns ➪ arrival in region after retreat of glacial ice either from Continental Europe or Volga-Ural region – haplogroup Z at low frequencies in Saami (Norw. Saami 0; Finnish Saami 7.2%; Northern Swedish Saami .7%; Southern Swedish Saami 4.3%) and Northern Asian population but virtually absent in Europe ➪ some Saami

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THOUGHTS ABOUT SAAMI PREHISTORY lineages shared common ancestor with lineages from Volga-Ural region as recently as 2700 BP. Johansson et al. 2004 “Linkage disequilibrium […]”: linkage disequilibrium affected by population demography: high LD in populations with relatively limited and constant size, because of genetic drift: Swedish Saami up to 6 Mb 3.61% vs. Swedish 35%, hence more LD. Kaessmann et al. 2002 “Extensive Linkage Disequilibrium […]”: LD – pairs in LD Swedish Saami 15 vs. Swedish 1, Finns 1 – increased genetic drift for Saami due to population size – Swedes and Finns descended from same central European source population – Finns’ higher LD vs. Swedes, .73 vs. .67 suggests founding population of Finns smaller than that of Swedes. Karlsson et al. 2006 “Y-chromosome diversity […]”: Y-chromosomal analysis in 7 Swedish regions, 1 Finnish region and a Swedish Saami population (Jokkmokk) – data indicate population continuity, acculturation and acceptance of new ideas rather than migration and population replacement – emerging agriculture an introduction of ideas. Kittles et al. 1998 “Dual Origins of Finns […]”: evidence of dual origins for Finns in pattern of Y chromosome variation in 9 Finnish provinces – two theories: 1) Finland founded ~2000 years ago by small number of settlers – then relative isolation, internal migration minimal until 17th, population growth not constant because of wars, famine, disease; 2) Finland settled by two different groups, first from the east near Lake Ladoga region, second from the south via Gulf of Finland – archaeological evidence supports dual origin model with considerable cultural differences between eastern and western Finland – Y chromosome haplotype A/49 common in northeast, haplotype B/69 common in south – A/49 predates B/69 by ~2000 years – B may have come in with group of agriculturists from the south. Lahermo et al. 1999 “Y chromosomal polymorphisms […]”: Y chromosome: C allele of Tat polymorphism common in Finno-Ugric: Finns 61.1%; Inari Saami 41.7%; Skolt Saami 61.7% – reduced variation (only a few Y chromosomal lineages in Finns and Saami) from bottlenecks – two major founding lineages in both Finns and Saami. Laitinen et al. 2002 “Y-chromosomal diversity […]”: Y-chromosomal variation in Estonian, Latvian and Lithuanian men suggests latter two are genetically similar to former – Baltic males share common Finno-Ugric-speaking forefathers. Meinilä et al. 2001 “Evidence for mtDNA Admixture […]”: high frequency of mtDNA haplotypes considered to be Saami specific in Finnish population suggest genetic admixture more pronounced in northern Finland – presence of haplogroup Z in Finns and Saami indicates that traces of Asian mtDNA have survived (4.3% of Saami and 3.9% of northern Finland. Raitio et al. 2001 “Y-Chromosomal SNPs […]”: two major SNP (single nucleotide polymorphism) haplotypes on Y chromosome in Finns and Saami – most common haplotypes are shared between Finns and Saami, but with regional differences ➪ supports hypothesis of two separate settlement waves to Finland (in accordance with archaeological data indicating dual origins for Finns) – haplotype H2 of Y-Chromosome DYS19 microsatellite locus: Finns 73% for allele 14, Saami (Kola, Skolt, Inari) 57% for allele 15;

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JOHN WEINSTOCK H16 DYS389-1 Saami 55% for allele 9, Finns 65% for allele 11, DYS 389-2 Saami 57% for allele 27, Finns 62% for allele 28. Richards et al. 1996 “Paleolithic and Neolithic Lineages […]”: major extant lineages throughout Europe predate Neolithic expansion and spread of agriculture was a substantially indigenous development accompanied by only a relatively minor component of contemporary Middle Eastern agriculturalists. Richards et al. 2000 “Tracing European Founder Lineages […]”: substantial backmigration into Near East; majority of extant mtDNA lineages entered Europe in several waves during Upper Paleolithic; founder effect or bottleneck associated with Last Glacial Maximum from which derive largest fraction of surviving lineages; immigrant Neolithic component less than ¼ of mtDNA pool of modern Europeans. Richards et al. 2002 “In Search of Geographical Patterns […]”: mtDNA exhibits significant geographical structuring. Rootsi et al. 2004 “Phylogeography […]”: analysis of HG I clade of Y phylogeny (I widespread in Europe but virtually absent elsewhere) – subclades of I have distinct geographic distributions with I1a mostly in Scandinavia. Ross et al. 2006 “Lifestyle, Genetics, and Disease in Sami”: heterogeneous genetic origin of Saami – authors survey origin of the Saami and suggest an area around Lake Ladoga and Lake Onega as a place of origin for the Saami. Rosser et al. 2000 “Y-Chromosomal Diversity […]”: Y-Chromosomal diversity in Europe is clinal, influence primarily by geography, rather than language. Sajantila 1995 “Mitochondrial DNA Diversity in Europe”: Saami (Finland, Sweden and Norway) show large amount of sequence differentiation compared to other European populations ➪ long history distinct from other populations. Sajantila et al. 1995 “Genes and Languages in Europe […]”: gene frequencies rather than genetic lineages reflect time perspectives more in concordance with linguistic evolution – Saami motif: Karasjok 52%; Skolt 36.2%; Norrbotten 32%; Inari 31.8%; Finn 2%; Karelian 6%. Sajantila et al. 1996 “Paternal and maternal DNA lineages […]”: Y-chromosomal haplotypes: near monomorphic pattern in Finns (94%, 2%), but less in Saami (68%, 11%) and Swedes (62%, 25%) + decrease in genetic diversity in mitochondrial control region ➪ few men and women have contributed to genetic lineages of Finns and a bottleneck ca. 4000 BP. Semino et al. 2000 “The Genetic Legacy […]”: Y chromosome perspective of Europe – ten lineages account for > 95% of Y chromosomes studied – geographic distribution and age compatible with two Paleolithic migratory episodes and one Neolithic migratory episode that have contributed to modern European gene pool – previously categorized Sardinians, Basques and Saami outliers share basically same Y binary components of other Europeans; their peculiar position probably consequence of genetic drift and isolation. Tambets et al. 2004 “The Western and Eastern […]”: origin of V and U5b1b is in western Europe; haplogroup H1, spread among the Saami, was absent in Samoyed and Ob-Ugric Siberians, but present in western and central European populations – best explanation for

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THOUGHTS ABOUT SAAMI PREHISTORY wide genetic separation of Saami from other Europeans is that the former are descendants of a narrow, distinctive subset of Europeans – extremes of genetic drift, such as due to repeated bottlenecks, could explain how the Saami mtDNA pool evolved as a narrow subset – Ychromosome haplogroups N3 and R1a (~60% of Saami) likely reached Fennoscandia from eastern Europe. Torroni et al. 1998 “mtDNA Analysis Reveals […]”: haplogroup V distribution: >40% of Saami have mtDNA haplogroup V, but low sequence variation and with only the two most common HVS-1 sequences ➪ Saami acquired V recently from other northern European populations and that it is associated with a strong founder event in the maternal lineage – V plus U make up more than 78% of Saami mtDNAs ➪ founder effects and genetic drift – less than 6.3% of Saami mtDNAs come from Asian superhaplogroup M – Y-chromosomal T ➝ C transition in some central-eastern Asian populations including Yakuts with 85.7%: Saami have 25% and Finns 52.4% ➪ Saami and Finns share part of their gene pool with western-central Asian populations. Torroni et al. 2001 “A Signal, from Human mtDNA […]”: haplogroup V arose in southwestern European refugium soon after late glacial maximum – distribution of V: Finns 2.5%; Skolt 52%; Inari 7.1%. Villems et al. 1998 “Reconstruction of Maternal lineages […]”: most Saami mtDNA display same basic European set of mtDNA varieties, also shared by Afro-Asiatic languagespeaking populations of Algeria, Middle East, Turks, i.e. Caucasoid – authors reject language-replacement model – Y chromosome papers: deletion mutant and T ➝ C transition are common in Saami and Finns but virtually absent from eastern Scandinavia to the west, though frequent in some Siberian populations – authors question “outlierness” of Saami and explain Y chromosome and mtDNA differences by drift. Zerjal et al. 1997 “Genetic Relationships of Asians […]”: T ➝ C transition likely occurred only once – probably in Asia – most of C-allele found among speakers of Altaic and Uralic – presence among Finns and Saami ➝ the chromosomes have been carried westward by migrations of Uralic-speaking populations – T ➝ C in Saami 9 to 3; in Finns 10 to 11.

Bibliography Appelbaum, Diana Muir & Paul S. Appelbaum. 2007. The gene wars. Azure 27 (5767 / 2007), pp. 1–13. Barth, Fredrik. 1969. Introduction. Ethnic Groups and Boundaries: The Social Organization of Cultural Difference, pp. 9–38. Ed. F. Barth. Boston: Little, Brown. Campbell, Lyle. 1997. On the linguistic prehistory of Finno-Ugric. Language History and Linguistic Modelling: A Festschrift for Jacek Fisiak on His 60th Birthday (Trends in Linguistics), pp. 829–862. Eds. Raymond Hickey & Stanislaw Puppel. Berlin: Mouton de Gruyter. Carpelan, Christian. 2000. Where do Finns come from?” http://sydaby.eget.net/swe/ jp_finns.htm (Available Febr. 11, 2009) Dolukhanov, Pavel M. 1996. The Early Slavs: Eastern Europe from the Initial Settlement to the Kievan Rus. London: Longman.

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JOHN WEINSTOCK Düben, Gustaf von. 1873. Om Lappland och lapparne, företrädesvis de svenske: etnografiska studier. Stockholm: Norstedt. Gray, Russell D. & Quentin D. Atkinson. 2003. Language-tree divergence times support the Anatolian theory of Indo-European origin. Nature 426, pp. 435–439. Hallström, Gustaf. 1929. Kan lapparnas invandringstid fixeras? Norrlands försvar 1929, pp. 39–92. Hansen, Lars Ivar & Bjørnar Olsen. (2004) 2007. Samenes historie: fram til 1750. Oslo: Cappelen. Kivikoski, Ella. 1961. Suomen esihistoria (Suomen historia 1). Ed. J. Jaakkola. Porvoo: WSOY. Manker, Ernst. 1952. Swedish contributions to Lapp ethnography. The Journal of the Royal Anthropological Institute of Great Britain and Ireland 82, 1, pp. 39–54. Nilsson, Sven. 1862–66. Skandinaviska nordens ur-invånare, ett försök i komparativa ethnografien och ett bidrag till menniskoslägtets utvecklings-historia. 1–2. Stockholm: Norstedt. Niskanen, Markku. 2002. The origin of the Baltic-Finns from the physical anthropological point of view. Mankind Quarterly 43, 2, pp. 121–153. Nummedal, Anders. 1929. Stone Age Finds in Finnmark. Oslo: Aschehoug. Nuñez, Milton. 2002. Finland’s settling model revisited. Mankind Quarterly 43, 2, pp. 155– 175. Renfrew, Colin. 1992. Archaeology, genetics and linguistic diversity. Man 27, pp. 445–478. — 2000. At the edge of knowability: towards a prehistory of languages. Cambridge Archaeological Journal 10, 1, pp. 7–34. Sammallahti, Pekka. 1998. The Saami Languages: An Introduction. Karasjok: Davvi Girji. — 1995. Language and roots. Orationes plenariae et conspectus quinquennales (Congressus Internationalis Fenno-Ugristarum, Jyväskylä 10.–15.8.1995, 1), pp. 143–155. [Ed.] H. Leskinen et al. Jyväskylä: Moderatores. Wiik, Kalevi. 2006. Who are the Finns? A Man of Measure: Festschrift in Honour of Fred Karlsson on his 60th Birthday (A Special Supplement to Journal of Linguistics 19), pp. 97–108. Turku: The Linguistic Association of Finland. Wiklund, K. B. 1947. Lapparna (Nordisk kultur 10). Stockholm: Bonniers. Zvelebil, M. & K. V. Zvelebil. 1988. Agricultural transition and Indo-European dispersals. Antiquity 62, pp. 574–578.

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